Wood-inhabiting Basidiomycetes in the Caucasus Region : Systematics and Biogeography

The Caucasus region is a hotspot of biodiversity and is one of the few areas in the Northern Hemisphere which harbor Pleistocene glacial refugia. The region encompasses Armenia, Azerbaijan, Georgia, the southernmost European Russia, NE Turkey, and northern Iran. The study on fungal composition of the Caucasus region and its connection and possible contribution to the present mycota of Europe has largely escaped empirical scrutiny. Using taxonomic surveys, phylogenetic reconstruction methods, haplotype analysis, and similarity tests, this study has aimed to, 1) summarize the knowledge on the occurrence of corticioids and polypores in the Caucasus region, 2) resolve the phylogenetic relationships of selected, resupinate wood-inhabiting basidiomycetes for which the Caucasus region is currently the mere, or one of the noteworthy areas of distribution, and, 3) assess the similarity of Caucasian corticioid fungi to those of Europe and important areas in the Northern Hemisphere, and to examine the significance of the Caucasus region as a glacial refugium for these fungi. This study provides the first catalogue of corticioids and polypores (635 species) occurring in the Caucasus region. The phylogeny and systematics of the Caucasian resupinate taxa in focus has been resolved and the usefulness of some morphological characters has been re-evaluated. In this context, four new genera and two new species were described and five new combinations were proposed, two of which were supplemented with modern descriptions. The species composition of corticioids in the Caucasus region is found to be distinctly more similar to Europe and North America than to East Asia and India. The highest molecular diversity and within population pairwise distance for Peniophorella praetermissa has been detected in the Caucasus and East Asia, with the isolates of the latter area being highly divergent from the European ones. This, and the assignment of root haplotype to the Caucasian isolates in a haplotype network for Phlebia tuberucalta and P. livida, call attention to the role of the Caucasus region in shaping the current mycota of Europe.

Phylogeography and demographic history of Lacerta lepida in the Iberian Peninsula

The Iberian Peninsula is recognized as an important refugial area for species survival and diversification during the climatic cycles of the Quaternary. Recent phylogeographic studies have revealed Iberia as a complex of multiple refugia. However, most of these studies have focused either on species with narrow distributions within the region or species groups that, although widely distributed, generally have a genetic structure that relates to pre-Quaternary cladogenetic events. In this study we undertake a detailed phylogeographic analysis of the lizard species, Lacerta lepida, whose distribution encompasses the entire Iberian Peninsula. We attempt to identify refugial areas, recolonization routes, zones of secondary contact and date demographic events within this species. Results support the existence of 6 evolutionary lineages (phylogroups) with a strong association between genetic variation and geography, suggesting a history of allopatric divergence in different refugia. Diversification within phylogroups is concordant with the onset of the Pleistocene climatic oscillations. The southern regions of several phylogroups show a high incidence of ancestral alleles in contrast with high incidence of recently derived alleles in northern regions. All phylogroups show signs of recent demographic and spatial expansions. We have further identified several zones of secondary contact, with divergent mitochondrial haplotypes occurring in narrow zones of sympatry. The concordant patterns of spatial and demographic expansions detected within phylogroups, together with the high incidence of ancestral haplotypes in southern regions of several phylogroups, suggests a pattern of contraction of populations into southern refugia during adverse climatic conditions from which subsequent northern expansions occurred. This study supports the emergent pattern of multiple refugia within Iberia but adds to it by identifying a pattern of refugia coincident with the southern distribution limits of individual evolutionary lineages. These areas are important in terms of long-term species persistence and therefore important areas for conservation.

Phylogeography and non-invasive landscape genetics of the European pine marten ("martes martes" L. 1758) : insights into ancient and contemporary processes shaping genetic variation

283 p. : graf., map.

An integrated assessment of the continued spread and potential impacts of the colonial ascidian, Didemnum sp. A, in U.S. waters

Didemnum sp. A is a colonial ascidian or “sea squirt” of unknown geographic origin. Colonies of Didemnum sp. A were first documented in U.S. waters in 1993 at Damariscotta River, Maine and San Francisco Bay, California. An alarming number of colonies have since been found at several locations in New England and along the West Coast of the contiguous continental United States. Originally believed to be restricted to artificial structures in nearshore habitats, such as ports and marinas, colonies of Didemnum sp. A have also been discovered on a gravel-pavement habitat on Georges Bank at depths of 40-65m. The wide distribution of Didemnum sp. A, the presence of colonies on an important offshore fishing ground, and the negative economic impacts that other species of noninidigenous ascidians have had on aquaculture operations have raised concerns about the potential impacts of Didemnum sp. A. We reviewed the available information on the biology and ecology of Didemnum sp. A and potentially closely related species to examine the environmental and socioeconomic factors that may have influenced the introduction, establishment and spread of Didemnum sp. A in U.S. waters, the potential impacts of this colonial ascidian on other organisms, aquaculture, and marine fisheries, and the possibility that it will spread to other U.S. waters. In addition, we present and discuss potential management objectives for minimizing the impacts and spread of Didemnum sp. A. Concern over the potential for Didemnum sp. A to become invasive stems from ecological traits that it shares with other invasive species, including the ability to overgrow benthic organisms, high reproductive and population growth rates, ability to spread by colony fragmentation, tolerance to a wide range of environmental conditions, apparent scarcity of predators, and the ability to survive in human dominated habitats. At relatively small spatial scales, species of Didemnum and other nonindigenous ascidians have been shown to alter the abundance and composition of benthic assemblages. In addition, the Canadian aquaculture industry has reported that heavy infestations of nonindigenous ascidians result in increased handling and processing costs. Offshore fisheries may also suffer where high densities of Didemnum sp. A may alter the access of commercially important fish species to critical spawning grounds, prey items, and refugia. Because colonial ascidian larvae remain viable for only 12–24hrs, the introduction and spread of Didemnum sp. A across large distances is thought to be predominantly human mediated; hull fouling, aquaculture, and ballast water. Recent studies suggest that colony growth rates decline when temperatures exceed 21 ºC for 7 consecutive days. Similarly, water temperatures above 8 to 10 ºC are necessary for colony growth; however, colonies can survive extended periods of time below this temperature threshold as an unidentified overwintering form. A qualitative analysis of monthly mean nearshore water temperatures suggest that new colonies of Didemnum will continue to be found in the Northeast U.S., California Current, and Gulf of Alaska LMEs. In contrast, water temperatures become less favorable for colony establishment in subarctic, subtropical, and tropical areas to the north and south of Didemnum’s current distribution in cool temperate habitats. We recommend that the Aquatic Nuisance Species Task Force serve as the central management authority to coordinate State and Federal management activities. Five objectives for a Didemnum sp. A management and control program focusing on preventing the spread of Didemnum sp. A to new areas and limiting the impacts of existing populations are discussed. Given the difficulty of eradicating large populations of Didemnum sp. A, developing strategies for limiting the access of Didemnum sp. A to transport vectors and locating newly established colonies are emphasized. (PDF contains 70 pages)

Untrawlable Bottom in Shrimp Statistical Zones of the Northwest Gulf of Mexico

The Gulf of Mexico Fisheries Management Council tasked the National Marine Fisheries Service with determining the extent, if any, of loss oft rawlable bottom in the Gulf of Mexico based upon fishing industry concerns. There are approximately 31 million hectares in the 21 shrimp statistical zones in the Gulf, approximately 23 million hectares of waters that are <35 fathoms (where most shrimp trawling effort occurs), and approximately 11 million hectares in zones 10-21, <35f athoms, which were examined. There are 31,338 known hangs, snags, artificial reefs, hazards to navigation, oil rigs, and similar obstructions which cause trawling to be unfeasible in these zones. There are several refuge (i.e. untrawlable) areas associated with the Alabama Artificial Reefs. Conservatively assuming 1 hectare for each known obstruction, coupled with the known area of each refuge, the estimate of total untrawlable bottom in zones 10-21 less than 35 fathoms in the Gulf is 185,953 hectares, or roughly 1.7% of this total trawlable area. Sensitivity analysis demonstrated the robustness of this assumption, with a range of 0.3-4.3% possible. In specific shrimp zones, untrawlable area is much less than 1% except in zones 10 (26%) and 11(2.5%), both of which possess a refuge. Other than the implementation periods of these refugia, no temporal trends were detectable with respect to the amount of untrawlable bottom.

根据特有植物的分布探讨中国冰期避难所

第三纪晚期和第四纪全球变冷对植物类群现在的分布式样有着重要的影响。植物特有现象出现的地区或中心一般都和一些适宜植物生存的避难所相联系，在这些避难所，众多植物类群得以保存、同时不断多样化形成新的类群。尽管中国区系中包括了丰富的特有类群，对特有类群丰富的地区也开展了一些初步的研究，但迄今我们对中国特有地区的数量、具体位置及其地理式样的了解仍然有限，对特有现象形成的原因更是所知甚少。为此，本研究通过选择555种中国特有植物类群（包括种、亚种和变种）为代表，对中国维管植物的特有中心及其地理式样进行了研究，并在此基础上探究了中国的冰期避难所。我们首先将这些特有类群的分布描绘在1  1纬度/经度的方格里，计算了特有现象的两个基本参数：总特有和加权特有，将那些特有参数值在前5%的单元确定为特有中心。 为了进一步了解中国的特有中心主要是植物的保存地（“植物博物馆”）还是植物新类群发生和物种形成（“植物摇篮”）的场所，或二者兼而有之，我们又根据化石证据、分子和形态系统树，以及生物地理等一系列数据，把555个特有类群分为古特有和新特有两个级别，探讨了这两个特有级别各自的分布式样。结果显示，具有特有现象的地区有20个，这些地区位于华中和华南，大致对应于这些地区的山系。特有中心主要位于青藏高原的东部（横断山和大雪山）、云贵高原、华中山脉、南岭山脉和东南山脉;海南和台湾也具有很多特有种。通过特有种所在的地区界定冰期避难所与第四纪植被重建的结果在很大程度上相一致，后者显示在这些地区在冰期有泛温带和亚热带森林出现。因此，山脉地区可能通过复杂的地形和更新世极端气候条件的局域缓冲为这些特有类群提供稳定生存的生境;这既保存了这些孑遗类群（“植物博物馆”），又使得新的类群得以不断出现（“植物摇篮”）。 中国特有中心既包含孑遗的类群（古特有）也包含近期新形成的类群（新特有）。然而，它们的分布和密度却非常不同，这可能与这些地区的地质分化和构造历史有关。尤其值得注意的是，青藏高原东部边缘是中国最重要的“进化前沿”，这可能与晚第三纪青藏高原的不断隆升有关。相反，在第三纪多数时候，华中和华南（除了海南和台湾）在地质构造上的相对稳定，使得孑遗植物谱系得以最大程度地保存。两种特有级别在全国水平上的比例（古特有为39.1%，新特有为60.9%）说明，尽管不同地区间存在一定的差异，但华中和华南既是晚第三纪和第四纪全球变冷后植物得以保存的避难所，也是植物分化和新类群形成的重要中心。 这些研究结果对特有植物、尤其对具有不同保护要求的孑遗类群和新类群的保护有着重要意义。我们可以针对富有古特有的地区以及有更多新类群产生的“植物摇篮”制定不同的保护策略和措施。

Changes in frost frequency and desert vegetation assemblages in Grand Canyon, Arizona

The effect of decreasing frost frequency on desert vegetation was documented in Grand Canyon by replication of historical photographs. Although views by numerous photographers of Grand Canyon have been examined, 400 Robert Brewster Stanton and Franklin A. Nims views taken in the winter of 1889-1890 provide the best information on recent plant distribution. In Grand Canyon, where grazing is limited by the rugged topography, vegetation dynamics are controlled by climate and by demographic processes such as seed productivity, recruitment, longevity and mortality. The replicated photographs show distribution and abundance of several species were limited by severe frost before 1889. Two of these, brittlebush (Encelia farinosa) and barrel cactus (Ferocactus cylindraceus), have clearly expanded their ranges up-canyon and have increased their densities at sites where they were present in 1890. In 1890, brittlebush was present in warm microhabitats that provided refugia from frost damage. Views showing desert vegetation in 1923 indicate that Encelia expanded rapidly to near its current distribution between 1890 and 1923, whereas the expansion of Ferocactus occurred more slowly. The higher frequency of frost was probably related to an anomalous increase in winter storms between 1878 (and possibly 1862) and 1891 in the southwestern United States.

Trends in counts of manatees Trichechus manatus latirostris from 1987 to 2006 in waters of Sarasota County, Florida, USA

To demonstrate the utility of distributional surveys for assessing relative abundance and trends in counts for a discrete area of coastline, aerial survey data from Sarasota County, Florida, USA, were analyzed for the years 1987 to 2006. The study area was divided into 3 regions: the Sarasota Bay Region (SBR; N = 353 surveys), Lemon Bay (N = 368), and the Myakka River (N = 209). Manatee counts varied significantly across seasons (p < 0.0001) for all 3 regions. Manatees within Sarasota County utilized open bays primarily in the warmer months. Such usage may have been influenced by resource availability. Conversely, usage of the Myakka River peaked in winter months when manatees seek warm-water refugia such as Warm Mineral Spring. Marginal means for yearly counts within Lemon Bay and the SBR increased significantly, beginning midway through the survey period (1996) until the early 2000s. In contrast, mean yearly counts within the Myakka River decreased over this time period. After record lows in 2003 for Lemon Bay and the Myakka River, and a considerable decline in 2004 for the SBR, mean yearly counts for all 3 regions showed an increasing trend over the remaining 2 yr of the study. Greater protection of manatee habitat and availability of forage coincided with the increase in numbers of manatees using Sarasota County waters during the 1990s, and the subsequent decline in numbers may be indicative of the increase in mortality in recent years due to watercraft collisions and severe red tide events.

Fish species diversity in the Victoria and Kyoga lake basins: their conservation and sustainable use

Introduction of exotic fish species especially the Nile perch Lates niloticus, is believed to be responsible for the decline of fish species diversity in lakes Victoria, Kyoga and Nabugabo.About 60% of the haplochromine cichlids are thought to have become extinct from L. Victoria due to predation by the Nile perch. However there are many lakes satelite to the lakes Victoria and Kyoga basins which still have fish fauna similar to that of the main lakes. many of the satellite lakes are separated from the main lakes in, which Nile perch was introduced by extensive swamps that provide a barrier to Nile perch .A survey was carried out in a number of these satelite lakes and an inventory made of existing fish species. Their distribution and relative abundances were also determined. The lakes studied included Nawampasa, Nakuwa,Kawi Lamwa Gigate, Nyaguo, Agu, Nabugabo. Kayanja, Kaytigi, Mburo, Kachera and Wamala.Some habitats within the main lakes Victoria and Kyoga, especially those with rocky outcrops· and macrophyte cover that provide refugia for endangered species from Nile perch,were also surveyed) Various stations along the River Nile were also sampled to quantify the fish species that are still resent. Kyoga minor lakes were found to have the highest number of fish species especially of haplochromine cichlids. Many haplochromine trophic groups that were thought to be extinct from 1. Victoria still occur in these lakes.!Some of the satellite lakes, especially lakes Kayugi, Mburo and Kachera still contain .healili populations of oreochromis. I esculentus that could be used as brood stock in fish farming. Many of these lakes should .I ( I therefore be protected for conservation offish species diversity

The biology, ecology, distribution and conservation of some surviving native non-cichlid fish species of Lakes Victoria, Kyoga and Nabugabo

Prior to introduction of non-native fish species into Lakes Victor i a, Kyoga and Nabugabo, the three lakes suppor ted diverse fish fauna representing 13 families consisting of six cichlid genera and fifteen non-cichlid genera. There were about 50 non-cichlid species and over 300 cichlids consisting of mainly haplochromines (Graham 1929, worthington 1929, Greenwood 1960). Many of the species were commercially and scientifically important and provided a rich variety of protein source to choose from. Following introduction of the Nile perch and several tilapiines species, most of the native species were drastically reduced and some have apparently disappeared. The few remaining species appear to be restricted in distribution due to the presence of the Nile perch. They are mainly confined to refugia such as marginal macrophytes, rocky outcrops and small satellite lakes which are separated from the areas of introduction by swamps

A report of the gillnet survey in the Ugandan waters of Lake Victoria under IFMP (January to March 2006)

Under the Implementation of the Fisheries Management Plan (IFMP) for Lake Victoria Result area 4, quarterly gillnet surveys are undertaken to monitor changes in fish stocks and environmental parameters in the shallow nontrawlable areas of the lake For purposes of monitoring surveys, the Ugandan sector of Lake Victoria is divided into 3 zones as shown in Figure 1. During the second quarter of APE2, two gillnet surveys were undertaken in zones 1 and 3 in February and March 2006 respectively. The purpose of the surveys was to monitor changes in the fish stocks and their biological characteristics, water quality, algal dynamics and invertebrate communities; as detailed in the various sections of the report. The surveys followed those conducted in November and December 2006 in the same zones. Results of the surveys showed that the number of fish taxa was higher in the near-shore fleets (0-100m) decreasing towards offshore. The near-shore areas were also associated with high primary productivity and hence secondary production to which Caridina and other invertebrates are part. These organisms are an important source of food for the fish and this may partly account for the high number of fish species recorded in this area of the lake. It was also observed that although Nile perch was the most dominant fish species recorded in all the stations during the surveys, haplochromines, Brycinus sadleri, Brycinus jacksonii Oreochromis niloticus and various mormyrid species contributed significantly to the fish biomass. The presence of many fish species and their coexistence with the predator, Nile perch is attributed to the presence of macrophyte cover and rocky habitats which serve as refugia in the shallow inshore habitats of Lake Victoria. In addition, the vegetated habitats are an important source of food for the fishes. As reported in macro-invertebrate studies, big populations of Caridina and other invertebrates were recorded among macrophyte beds. Caridina is an important source of food for juvenile Nile perch and other fish species so are the other invertebrates especially chironomid larvae, odonata nymphs and molluscs. Resurgence in haplochromine cichlids was observed during the surveys. The presence of haplochromines cichlids in all the sites especially Thruston Bay where it ranked the second by percentage contribution in number, is evidence of the recovery of this group of fishes which had declined largely due to predation by L. niloticus. Caridina nilotica has also increased in biomass and is a major component of the Nile perch diet. This could have reduced predation pressure on the haplochromines by Nile perch and has possibly contributed to recent resurgence in haplochromines cichlids in the lake in the shallow nontrawlable areas of the lake Rastrineobola argentea was found to be an important prey item for Nile perch and other fish species such as Clarias gariepinus. Measures should therefore be taken to ensure sustainable harvesting of Dagaa so that there is enough left to sustain the fishery of Nile perch and other species.

The role of satellite lakes in conservation of fish species diversity in the Lake Kyoga basin

Lakes Victoria and Kyoga had, a diverse fish fauna, which was important as food for local population and valuable in scientific studies. Over the past twenty years, the diversity of fish in these lakes had declined due to over-exploitation, introduction of new fish species including the piscivorous Nile perch and degradation of fish habitat. Studies of satellite lakes in the Victoria and Kyoga lake basins suggested that some of these lakes harboured species which had been lost from the main lakes. In order to better understand the extent, to which these satellite lakes may serve as refugia, a faunal survey was undertaken to determine the distribution and nature of the taxa found. Seven satellite lakes and the eastern end of the main Lake Kyoga adjacent to these minor lakes were surveyed over a two-year period for fish species diversity. A total of 68 fish species were recorded of which 41 were haplochromines. Almost all the native non cichlids which occurred in the main lakes (Victoria and Kyoga) before the Nile perch upsurge recorded. Lakes Nawampasa, Gigati, Kawi, Agu and Nyaguo had the highest fish species and trophic diversity. The trophic diversity of haplochromines (based on Shannon Weaver Index) was highest in Lake Nawampasa (1.28), followed by Gigati (1.25), Kawi (1.18), Agu (0.8), Lemwa (0.81), Nyaguo (0.35) and was lowest in the main Lake Kyoga. Potential threats to these lakes were from collectors of ornamental fish species, especially the haplochromines, the spread of the predatory Nile perch and the water hyacinth, which are already in Lake Kyoga, and the destruction of macrophytes through harvesting of papyrus and reclamation for agriculture. The human population around these lakes harvested the fishes for food but the levels of exploitation were still low because the lakes were adjacent to main Lake Kyoga, the major supply of fish. Ornamental fish dealers were encouraged to start captive breeding of the fish for export to reduce pressure on the lakes and demonstrations for breeding were set up at FIRI in Jinja. Meetings and seminars were held with some of the communities living around the lakes sampled and the importance of fish species found in these lakes and the dangers of destructive practices discussed. Representatives of all taxa of fish caught from the lakes were preserved, catalogued and stored in the FIRI Museum. Results from this survey support the motion that these satellite lakes are important refugia for endemic diversity. Based on survey, we recommend that SaIne of these lakes like Nawampasa, Gigati, Kawi, Agu and Nyaguo could be designated as conservation areas of species threatened in the main lakes. One of the factors that seem to have prevented the spread of Nile perch into Kyoga Minor lakes seems to have been the presence of extensive swamps around these lakes and the low oxygen levels that exist in these habitats. Clearing of swamps and vegetation that separate Kyoga minor lakes from the main lake should be avoided to prevent Nile perch from spreading into these lakes.

PHYLOGEOGRAPHIC PATTERNS INDICATE TRANSATLANTIC MIGRATION FROM EUROPE TO NORTH AMERICA IN THE RED SEAWEED CHONDRUS CRISPUS (GIGARTINALES, RHODOPHYTA)1

Inferring how the Pleistocene climate oscillations have repopulated the extant population structure of Chondrus crispus Stackh. in the North Atlantic Ocean is important both for our understanding of the glacial episode promoting diversification and for the conservation and development of marine organisms. C. crispus is an ecologically and commercially important red seaweed with broad distributions in the North Atlantic. Here, we employed both partial mtDNA Cox1 and nrDNA internal transcribed spacer region 2 (ITS2) sequences to explore the genetic structure of 17 C. crispus populations from this area. Twenty-eight and 30 haplotypes were inferred from these two markers, respectively. Analysis of molecular variance (AMOVA) and of the population statistic Theta(ST) not only revealed significant genetic structure within C. crispus populations but also detected significant levels of genetic subdivision among and within populations in the North Atlantic. On the basis of high haplotype diversity and the presence of endemic haplotypes, we postulate that C. crispus had survived in Pleistocene glacial refugia in the northeast Atlantic, such as the English Channel and the northwestern Iberian Peninsula. We also hypothesize that C. crispus from the English Channel refugium repopulated most of northeastern Europe and recolonized northeastern North America in the Late Pleistocene. The observed phylogeographic pattern of C. crispus populations is in agreement with a scenario in which severe Quaternary glaciations influenced the genetic structure of North Atlantic marine organisms with contiguous population expansion and locally restricted gene flow coupled with a transatlantic dispersal in the Late Pleistocene.

Phylogeography of Pinus tabulaeformis Carr. (Pinaceae), a dominant species of coniferous forest in northern China

How coniferous trees in northern China changed their distribution ranges in response to Quaternary climatic oscillations remains largely unknown. Here we report a study of the phylogeography of Pinus tabulaeformis, an endemic and dominant species of coniferous forest in northern China. We examined sequence variation of maternally inherited, seed-dispersed mitochondrial DNA (mtDNA) (nad5 intron 1 and nad4/3-4) and paternally inherited, pollen- and seed-dispersed chloroplast DNA (cpDNA) (rpl16 and trnS-trnG) within and among 30 natural populations across the entire range of the species. Six mitotypes and five chlorotypes were recovered among 291 trees surveyed. Population divergence was high for mtDNA variation (G(ST) = 0.738, N-ST = 0.771) indicating low levels of seed-based gene flow and significant phylogeographical structure (N-ST > G(ST), P < 0.05). The spatial distribution of mitotypes suggests that five distinct population groups exist in the species: one in the west comprising seven populations, a second with a north-central distribution comprising 15 populations, a third with a southern and easterly distribution comprising five populations, a fourth comprising one central and one western population, and a fifth comprising a single population located in the north-central part of the species' range. Each group apart from the fourth group is characterized by a distinct mitotype, with other mitotypes, if present, occurring at low frequency. It is suggested, therefore, that most members of each group apart from Group 4 are derived from ancestors that occupied different isolated refugia in a previous period of range fragmentation of the species, possibly at the time of the Last Glacial Maximum. Possible locations for these refugia are suggested. A comparison of mitotype diversity between northern and southern subgroups within the north-central group of populations (Group 2) showed much greater uniformity in the northern part of the range both within and between populations. This could indicate a northward migration of the species from a southern refugium in this region during the postglacial period, although alternative explanations cannot be ruled out. Two chlorotypes were distributed across the geographical range of the species, resulting in lower levels of among-population chlorotype variation. The geographical pattern of variation for all five chlorotypes provided some indication of the species surviving past glaciations in more than one refugium, although differentiation was much less marked, presumably due to the greater dispersal of cpDNA via pollen.

Mitochondrial and chloroplast phylogeography of Picea crassifolia Kom. (Pinaceae) in the Qinghai-Tibetan Plateau and adjacent highlands

The disjunct distribution of forests in the Qinghai-Tibetan Plateau (QTP) and adjacent Helan Shan and Daqing Shan highlands provides an excellent model to examine vegetation shifts, glacial refugia and gene flow of key species in this complex landscape region in response to past climatic oscillations and human disturbance. In this study, we examined maternally inherited mitochondrial DNA (nad1 intron b/c and nad5 intron 1) and paternally inherited chloroplast DNA (trnC-trnD) sequence variation within a dominant forest species, Picea crassifolia Kom. We recovered nine mitotypes and two chlorotypes in a survey of 442 individuals from 32 populations sampled throughout the species' range. Significant mitochondrial DNA population subdivision was detected (G(ST) = 0.512; N-ST = 0.679), suggesting low levels of recurrent gene flow through seeds among populations and significant phylogeographical structure (N-ST > GST, P < 0.05). Plateau haplotypes differed in sequence from those in the adjacent highlands, suggesting a long period of allopatric fragmentation between the species in the two regions and the presence of independent refugia in each region during Quaternary glaciations. On the QTP platform, all but one of the disjunct populations surveyed were fixed for the same mitotype, while most populations at the plateau edge contained more than one haplotype with the mitotype that was fixed in plateau platform populations always present at high frequency. This distribution pattern suggests that present-day disjunct populations on the QTP platform experienced a common recolonization history. The same phylogeographical pattern, however, was not detected for paternally inherited chloroplast DNA haplotypes. Two chlorotypes were distributed throughout the range of the species with little geographical population differentiation (G(ST) = N-ST = 0.093). This provides evidence for highly efficient pollen-mediated gene flow among isolated forest patches, both within and between the QTP and adjacent highland populations. A lack of isolation to pollen-mediated gene flow between forests on the QTP and adjacent highlands is surprising given that the Tengger Desert has been a geographical barrier between these two regions for approximately the last 1.8 million years.